Nienke Hoeksema


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  • Hoeksema, N., Hagoort, P., & Vernes, S. C. (2022). Piecing together the building blocks of the vocal learning bat brain. In A. Ravignani, R. Asano, D. Valente, F. Ferretti, S. Hartmann, M. Hayashi, Y. Jadoul, M. Martins, Y. Oseki, E. D. Rodrigues, O. Vasileva, & S. Wacewicz (Eds.), The evolution of language: Proceedings of the Joint Conference on Language Evolution (JCoLE) (pp. 294-296). Nijmegen: Joint Conference on Language Evolution (JCoLE).
  • Vernes, S. C., Devanna, P., Hörpel, S. G., Alvarez van Tussenbroek, I., Firzlaff, U., Hagoort, P., Hiller, M., Hoeksema, N., Hughes, G. M., Lavrichenko, K., Mengede, J., Morales, A. E., & Wiesmann, M. (2022). The pale spear‐nosed bat: A neuromolecular and transgenic model for vocal learning. Annals of the New York Academy of Sciences, 1517, 125-142. doi:10.1111/nyas.14884.


    Vocal learning, the ability to produce modified vocalizations via learning from acoustic signals, is a key trait in the evolution of speech. While extensively studied in songbirds, mammalian models for vocal learning are rare. Bats present a promising study system given their gregarious natures, small size, and the ability of some species to be maintained in captive colonies. We utilize the pale spear-nosed bat (Phyllostomus discolor) and report advances in establishing this species as a tractable model for understanding vocal learning. We have taken an interdisciplinary approach, aiming to provide an integrated understanding across genomics (Part I), neurobiology (Part II), and transgenics (Part III). In Part I, we generated new, high-quality genome annotations of coding genes and noncoding microRNAs to facilitate functional and evolutionary studies. In Part II, we traced connections between auditory-related brain regions and reported neuroimaging to explore the structure of the brain and gene expression patterns to highlight brain regions. In Part III, we created the first successful transgenic bats by manipulating the expression of FoxP2, a speech-related gene. These interdisciplinary approaches are facilitating a mechanistic and evolutionary understanding of mammalian vocal learning and can also contribute to other areas of investigation that utilize P. discolor or bats as study species.

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  • Hoeksema, N., Verga, L., Mengede, J., Van Roessel, C., Villanueva, S., Salazar-Casals, A., Rubio-Garcia, A., Curcic-Blake, B., Vernes, S. C., & Ravignani, A. (2021). Neuroanatomy of the grey seal brain: Bringing pinnipeds into the neurobiological study of vocal learning. Philosophical Transactions of the Royal Society of London, Series B: Biological Sciences, 376: 20200252. doi:10.1098/rstb.2020.0252.


    Comparative studies of vocal learning and vocal non-learning animals can increase our understanding of the neurobiology and evolution of vocal learning and human speech. Mammalian vocal learning is understudied: most research has either focused on vocal learning in songbirds or its absence in non-human primates. Here we focus on a highly promising model species for the neurobiology of vocal learning: grey seals. We provide a neuroanatomical atlas (based on dissected brain slices and magnetic resonance images), a labelled MRI template, a 3D model with volumetric measurements of brain regions, and histological cortical stainings. Four main features of the grey seal brain stand out. (1) It is relatively big and highly convoluted. (2) It hosts a relatively large temporal lobe and cerebellum, structures which could support developed timing abilities and acoustic processing. (3) The cortex is similar to humans in thickness and shows the expected six-layered mammalian structure. (4) Expression of FoxP2 - a gene involved in vocal learning and spoken language - is present in deeper layers of the cortex. Our results could facilitate future studies targeting the neural and genetic underpinnings of mammalian vocal learning, thus bridging the research gap from songbirds to humans and non-human primates.Competing Interest StatementThe authors have declared no competing interest.
  • Gilbers, S., Hoeksema, N., De Bot, K., & Lowie, W. (2020). Regional variation in West and East Coast African-American English prosody and rap flows. Language and Speech, 63(4), 713-745. doi:10.1177/0023830919881479.


    Regional variation in African-American English (AAE) is especially salient to its speakers involved with hip-hop culture, as hip-hop assigns great importance to regional identity and regional accents are a key means of expressing regional identity. However, little is known about AAE regional variation regarding prosodic rhythm and melody. In hip-hop music, regional variation can also be observed, with different regions’ rap performances being characterized by distinct “flows” (i.e., rhythmic and melodic delivery), an observation which has not been quantitatively investigated yet. This study concerns regional variation in AAE speech and rap, specifically regarding the United States’ East and West Coasts. It investigates how East Coast and West Coast AAE prosody are distinct, how East Coast and West Coast rap flows differ, and whether the two domains follow a similar pattern: more rhythmic and melodic variation on the West Coast compared to the East Coast for both speech and rap. To this end, free speech and rap recordings of 16 prominent African-American members of the East Coast and West Coast hip-hop communities were phonetically analyzed regarding rhythm (e.g., syllable isochrony and musical timing) and melody (i.e., pitch fluctuation) using a combination of existing and novel methodological approaches. The results mostly confirm the hypotheses that East Coast AAE speech and rap are less rhythmically diverse and more monotone than West Coast AAE speech and rap, respectively. They also show that regional variation in AAE prosody and rap flows pattern in similar ways, suggesting a connection between rhythm and melody in language and music.
  • Hoeksema, N., Villanueva, S., Mengede, J., Salazar-Casals, A., Rubio-García, A., Curcic-Blake, B., Vernes, S. C., & Ravignani, A. (2020). Neuroanatomy of the grey seal brain: Bringing pinnipeds into the neurobiological study of vocal learning. In A. Ravignani, C. Barbieri, M. Flaherty, Y. Jadoul, E. Lattenkamp, H. Little, M. Martins, K. Mudd, & T. Verhoef (Eds.), The Evolution of Language: Proceedings of the 13th International Conference (Evolang13) (pp. 162-164). Nijmegen: The Evolution of Language Conferences.
  • Hoeksema, N., Wiesmann, M., Kiliaan, A., Hagoort, P., & Vernes, S. C. (2020). Bats and the comparative neurobiology of vocal learning. In A. Ravignani, C. Barbieri, M. Flaherty, Y. Jadoul, E. Lattenkamp, H. Little, M. Martins, K. Mudd, & T. Verhoef (Eds.), The Evolution of Language: Proceedings of the 13th International Conference (Evolang13) (pp. 165-167). Nijmegen: The Evolution of Language Conferences.
  • Ravignani, A., Garcia, M., Gross, S., de Reus, K., Hoeksema, N., Rubio-Garcia, A., & de Boer, B. (2018). Pinnipeds have something to say about speech and rhythm. In C. Cuskley, M. Flaherty, H. Little, L. McCrohon, A. Ravignani, & T. Verhoef (Eds.), Proceedings of the 12th International Conference on the Evolution of Language (EVOLANG XII) (pp. 399-401). Toruń, Poland: NCU Press. doi:10.12775/3991-1.095.

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